The Eco-Ethology of the Karoo Korhaan Eupodotis Virgorsil
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THE ECO-ETHOLOGY OF THE KAROO KORHAAN EUPODOTIS VIGORSII. BY M.G.BOOBYER University of Cape Town SUBMITIED IN PARTIAL FULFILMENT OF THE DEGREE OF MASTER OF SCIENCE (ORNITHOLOGY) UNIVERSITY OF CAPE TOWN RONDEBOSCH 7700 CAPE TOWN The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town University of Cape Town PREFACE The study of the Karoo Korhaan allowed me a far broader insight in to the Karoo than would otherwise have been possible. The vast openness of the Karoo is a monotony to those who have not stopped and looked. Many people were instrumental in not only encouraging me to stop and look but also in teaching me to see. The farmers on whose land I worked are to be applauded for their unquestioning approval of my activities and general enthusiasm for studies concerning the veld and I am particularly grateful to Mnr. and Mev. Obermayer (Hebron/Merino), Mnr. and Mev. Steenkamp (Inverdoorn), Mnr. Bothma (Excelsior) and Mnr. Van der Merwe. Alwyn and Joan Pienaar of Bokvlei have my deepest gratitude for their generous hospitality and firm friendship. Richard and Sue Dean were a constant source of inspiration throughout the study and their diligence and enthusiasm in the field is an example to us all. Many people were indispensible in showing me around the flora of the Karoo. Of these, Sue Dean, Craig Hilton-Taylor, Timm Hoffman, Bruce Bayer, Richard Cowling and David Shearing deserve my full gratitude as, were it not for each one of them .I wouldn't have made any headway at all. Graham Kerley who accompanied me on my first sojourn to the Karoo was always a source of enthusiasm and companionship. My friends Grant Fabian and Wayne Smith both gave indispensible help in the field, as did Jimmy McCrindle, Ali Mohammed and Dave Allan. Peter Steyn and Richard Brooke provided valuable accounts of their observations of korhaans. Of all the people who contributed to this study, Geoff Hoy stands out as an uncomplaining technical advisor and close friend whose skill and encouragement were indispensible and kept me going throughout. Many people provided valuable assistance during the process of printing the final draft. Of these, Mark Gibbons, Elaine Rumbak, Morne Du Plessis and Ken Findlay deserve special thanks. Mark Gibbons was partly responsible for my original decision to move to South Africa to study and for his support as a friend and his criticism as a scientist I will always be grateful. Geordie Ractliffe provided support during the most trying times and her companionship gave me strength when it was most needed. I will alwaysUniversity be in the debt of the University of Capeof Cape Town andTown the Council for Scientific and Industrial Research who jointly supported me for the first two years of this study. The FitzPatrick Institute of African Ornithology was a sound base from which to project myself into the Karoo and the staff and facilities of the "Fitz" were, in no small way, really responsible for this project being possible. My thanks are deserved by my supervisor, Dr. Phillip Hockey, who provided valuable criticism of this manuscript. Finally, I thank my family who have had no experience of the large, dry, open spaces of the Karoo, but who have patiently indulged my absence and encouraged me throughout. I dedicate this work to them. TABLE OF CONTENTS Chapter One 1 General introduction 1 Chapter Two 4 Pbytosociological classification of Karoo Korhaan habitats in the Succulent and Nama-Karoo biomes. 4 Introduction 4 Methods 4 Results 5 Touwsrivier Succulent Karoo. 5 Study area 5 Results 5 Discussion 6 Ceres Succulent Karoo 7 Study area 7 Results 8 Discussion 9 Western Mountain Karoo 9 Study area 9 Results 10 Discussion 10 The Great Karoo 11 Study area 11 Results 11 Discussion 12 General conclusions 12 Chapter Three 14 Physiognomic classification of Karoo Korhaan habitats in the Succulent and Nama-Karoo biomes. 14 Introduction 14 Study areas 14 Methods 14 Results 15 Spring 15 Summer 16 Autumn 16 Winter 17 General results 17 Discussion University of Cape Town 18 Chapter Four 21 The distribution of Karoo Korhaans in relation to the phytosociology and physiognomy of karroid habitats 21 Introduction 21 Methods 21 Results 21 Phytosodology 21 Discussion 23 Chapter Five 25 Diet of the karoo Korhaan 25 Introduction 25 Methods 25 Presentation of the results 27 Results 29 Oesophagus and proventriculus 29 Conclusions of the analysis of the contents of the proventriculus and oesophagus.' 29 Analysis of the stomach contents 30 Animal material 30 Plant material 33 General results 36 Discussion 38 Chapter Six 42 Group size in the Karoo Korhaan 42 Introduction 42 Methods 42 ~~ ~ Behavioural observations of Karoo Korhaans - defending territories 44 Group sizes 44 Discussion 45 Chapter Seven 49 Is the Karoo Korhaan an indicator species? 49 Introduction 49 Results 50 Conclusions 51 Chapter Eight 53 Conclusions 53 References 56 Appendix one Information statistics tables 66 Appendix two Box and whisker plots 75 Appendix three Denizens of the desert 82 University of Cape Town · CHAPTER ONE GENERAL INTRODUCTION University of Cape Town 1 The following study was undertaken as part of the Karoo Biome Project (KBP) and sets out to investigate an aspect of karoo ecology that will contribute to a predictive understanding of both biotic and abiotic processes in the region for the development of meaningful management guidelines (Cowling, 1986). The Karoo is a vast arid to semi-arid region south of 22°S, covering 652 339 km2 of southern Africa (Rutherford & Westfall, 1986). In South Africa alone, the Karoo accounts for 427 015 km2 or 35% of the land surface area (Cowling, 1986). The karoo biome (sensu Huntley, 1974) is an area of extensive pastoralism, the wide ranging non-selective grazing of large migratory herbivores (most typically SpringbokAntidorcas marsupialis) having been replaced by a profitable small stock industry. The generally lime-rich, weakly developed soils are readily erodable, and, in areas where persistent overgrazing has significantly reduced the natural vegetation cover, gully and sheet erosion have reached advanced stages. Typified by extensive flat to gently undulating plains with characteristic dolerite topped "Mesas" and some mountainous regions, the area ranges from sea level in some western areas to 2000 m above sea level in the interior. Arid and semi-arid areas generally are defined as areas where potential evaporation exceeds the average precipitation, and are characterized by rainfall that is not only limited, but also extremely irregular, both in timing and the amount it eventually contributes to soil moisture (Noy-Meir, 1973; Mott, 1979). The Karoo is particularly unfavourably placed geographically relative to the rain bearing circulation patterns of southern Africa (Booysen & Rowswell, 1983). The sub-continent is situated largely within the southern sub-tropical high pressure belt and is skirted to the south by circumpolar westerly airstreams (Weather Bureau, 1950). Rainfall decreases to the west and to the north of the region (Anon, 1957) ranging from 20-290mm in the north-west to 100-520mm in the east (Rutherford & Westfall, 1986). Linked with a decrease in the annual precipitation is a decrease in its reliability. The relative variability of the rainfall is about 25-30%, increasing to 80% in the north-west (Werger, 1978). Precipitation in the summer rainfall areas of the interior of South Africa originates mostly from the extension of thunderUniversity storm activity emanating fromof theCape north and east. Town The western regions receive up to 60% of their precipitation from the deep penetration of cyclonic fronts cut off from the low pressure cell in spring and autumn (Cowling et. al., 1986). Although generally divided into summer and winter rainfall regions the whole area experiences summer drought (Cowling, 1987). A more comprehensive review of the physical environment of the karoo biome is provided by Cowling et. al., 1987a,b. The flora of the Karoo is dominated by members of the Asteraceae, Mesembryanthemaceae, Aizoaceae, Poaceae and Scrophulariaceae (Werger, 1978; Hilton-Taylor, 1987). There is an increasing dominance of dwarf and low succulent shrubs towards the western and southern continental margins associated with a decrease in the likelihood of severe winter frost in these regions (Werger, 1978). Rutherford and Westfall's (1986) separation of the region into two "biomes" on the basis of floral life forms, an index of summer aridity and moisture regime places emphasis on existing 2 vegetation and not on supposed climax vegetation and productivity (Acocks, 1975). However, Acocks' (1975) categorization of the vegetation of South Africa remains the most detailed for the region and I refer to his "veld types" throughout. Rutherford and Westfall (1986) divide the area in to the Succulent Karoo biome (limited to even, winter and strong winter rainfall with the greatest summer aridity in South Africa) and the Nama-Karoo biome (limited to strong summer, summer and even rainfall). For a detailed description of these biomes and the criteria for their separation see Rutherford & Westfall (1986). Soil water in arid and semi-arid regions is far from being a single homogenous resource and the availability of soil moisture to different plant growth forms varies according to a suite of physiognomic and edaphic factors. In the summer rainfall areas a large proportion of the moisture does not penetrate beyond the grass rooting zone and therefore these areas are dominated by grasses (Roux, 1966; Werger, 1978).